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Incidence, Size and Spatial Structure of Clones in Second-Growth Stands of Coast Redwood, Sequoia Sempervirens (Cupressaceae)1

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Incidence, Size and Spatial Structure of Clones in Second-Growth Stands of Coast Redwood, Sequoia Sempervirens (Cupressaceae)1 American Journal of Botany 91(7): 1140±1146. 2004. INCIDENCE, SIZE AND SPATIAL STRUCTURE OF CLONES IN SECOND-GROWTH STANDS OF COAST REDWOOD, SEQUOIA SEMPERVIRENS (CUPRESSACEAE)1 VLADIMIR DOUHOVNIKOFF,2 ADELAIDE M. CHENG, AND RICHARD S. DODD Department of Environmental Science, Policy and Management, University of California, Berkeley, California 94720-3112 USA The ecology and evolutionary potential of coast redwood (Sequoia sempervirens) is signi®cantly in¯uenced by the important role clonal spread plays in its reproduction and site persistence. In nine second-growth stands, ampli®ed fragment length polymorphisms (AFLPs) were used to identify redwood clonal architecture. Clones (multistem genets) dominated sites by representing an average of 70% of stems measured, ranging in size from two to 20 stems. As a result, a relatively small number of genets can monopolize a disproportionate amount of site resources, are more likely to persist over time, and have greater on-site genetic representation. Clones were not limited to fairy-ring structures, but consisted of a wide range of shapes including concentric rings, ring chains, disjunct, and linear structures. Between-ramet distances of up to 40 m were measured, indicating that clonal reproduction is not limited to basal stump resprouting. Clonal structure in second-growth stands was similar to earlier reports from old growth, emphasizing the importance of site persistence and long-term, gradual site development. Smaller ramet numbers per genet in old growth is probably due to local within-genet self thinning. Management and conservation of redwoods will bene®t from a better understanding of the dynamics and structure of clonal spread in these forests. Key words: AFLPs; clonal spread; genet; molecular markers; ramet; Sequoia; spatial structure; woody plant. Asexual regeneration through sprouting from vegetative ma- son and Jerling, 1990). Evaluated ecologically, these bene®ts terial is a common phenomenon in plants (Stuefer et al., 2002), can result in high levels of productivity, in competitive ad- and many community types are dominated by such clonal spe- vantage for site colonization and persistence, and in an ability cies (van Groenendael and de Kroon, 1990; KlimesÏ et al., to survive a wide range of disturbances. 1997; Oborny and Kun, 2002). Commonly, clonal plants re- Through its in¯uence on plant ®tness, clonal spread also has produce both sexually, by the production and germination of important evolutionary implications. Clonal spread is the veg- seeds, and asexually, by sprouting from roots and shoots. De- etative establishment of genetically identical ramets that allow spite the fact that both means of reproduction play an impor- the persistence of a genet long after the original ortet is gone. tant role in the ecology and evolution of plant species (de There may be no limit to the number of ramet generations that Kroon and van Groenendael, 1990; Fischer and van Kleunen, can be generated over time within a clone, and each additional 2002; Pan and Price, 2002), the majority of research involving ramet adds to the overall ®tness of the genet through survi- plant population dynamics has focused on issues of sexual vorship, future seed production, and subsequent ramet produc- reproduction (fertilization, seed dormancy, longevity, morpho- tion (Fischer and van Kleunen, 2002; Pan and Price, 2002). logical characteristics, dispersal and germination requirements, In this way, clonal spread can in¯uence within- and among- etc.). This is re¯ected in the emphasis such reproduction is population genetic structure, effective population size, meta- given in most ecological and evolutionary theories and mod- population dynamics, natural selection, and possibly even the els. Yet, in many species the persistence of existing plant ge- evolution of geographic range (Eckert, 2002). notypes through clonal spread can be the more important av- Despite the potential importance of clonality and the fact enue to plant population regeneration and maintenance (Bond that many woody plants are capable of vegetative regeneration and Midgely, 2001). and clonal spread through sprouting (Peterson and Jones, The ecological signi®cance of clonal spread arises from the 1997), surprisingly few clonal studies have focused on trees bene®ts it confers to the plant. Some bene®ts include effective (Barsoum, 2002), and sprouting ``has received very little at- resource acquisition as a result of foraging (Oborny and Cain, tention in models and theories of forest succession or forest 1997), division of labor (Alpert and Stuefer, 1997), and re- diversity'' (Bond and Midgley, 2001, p. 49). Among woody source sharing (JoÂnsdoÂttir and Watson, 1997). Others include plants, the little work that has been reported has focused pri- better survival relative to seedlings of offspring in new and marily on angiosperms. sometimes dif®cult environments (Peterson and Jones, 1997), Redwoods (Sequoia sempervirens (D. Don) Endl.) are one reduced population turnover and dependence on seeds (Bond of the most productive timber types of the United States (Ol- and Midgley, 2001), and risk evasion on a genet scale (Eriks- iver et al., 1994) and provide an important supply of high- quality wood ®ber. Yet, beyond the fact that coast redwood is 1 Manuscript received 27 October 2003; revision accepted 16 March 2004. exceptional for a gymnosperm in its ability to readily resprout The authors thank Joe R. McBride and William L. Libby for their diverse and grow clonally after disturbances such as ®res, ¯oods, land- and important contributions, and William Baxter, Kevin L. O'Hara, Scott Ste- slides, and harvesting (Rydelius and Libby, 1993; Rogers, phens, John Battles, Matteo Garbelotto, and Nicholas Douhovnikoff for lab- oratory, ®eld, data, and manuscript assistance. This work was supported by a 1994, 2000), very little is known about the spatial structure of grant from the California Department of Forestry and Fire Protection. redwood clones, the relative importance of clonal vs. nonclon- 2 E-mail: [email protected]. al reproduction, or the variables that in¯uence clonal spread. 1140 July 2004] DOUHOVNIKOFF ET AL.ÐCLONAL SPREAD IN COAST REDWOODS 1141 Fig. 1. Sample sites of Sequoia sempervirens in the Jackson Demonstration State Forest, north coast of California, USA. Sites are represented by stars (river 5 R1, R2, R3; north-facing 5 N1, N2, N3; south-facing 5 S1, S2, S3) and rivers are marked as dotted lines. In a study of old-growth stands using allozyme markers, Rog- stands and explore the local structure of redwood clones. If ers (2000) found that a signi®cant number of redwood stems clonal spread is important in redwood stands, Bellingham and were not genetically distinct individuals, but were ramets of Sparrow (2000) predict that it should be in¯uenced by distur- larger genets, and thus resulted from cloning and not from bance frequency and site productivity. Therefore, we were also reproduction by seed. There are no similar studies of second- interested in determining if variability in broad site character- growth redwood forests despite the fact that in 1990 as much istics (i.e., aspect, disturbance levels) affects the importance as 96% of those lands in commercial redwood production of cloning and clone structure at a stand scale. available for harvest were made up of young or second-growth stands (California Department of Forestry and Fire Protection, MATERIALS AND METHODS 1990). If we are to properly understand the ecology and evolution- The study was conducted in the Jackson Demonstration State Forest (Fig. ary potential of redwoods, and clonal woody plants in general, 1) because of its combination of historic intensive management and the avail- it is essential to further explore the structure, dynamics, and ability of mature second-growth stands. A total of nine sites were selected (Table 1). Three sites each were selected on north-facing slopes (N), south- signi®cance of clonal spread in these forests. In the context of facing slopes (S), and on ¯oodplains adjacent to rivers (R). These broad cat- management and conservation, for example, this would result egories were chosen in order to investigate possible differences resulting from in more accurate estimates of minimum viable population size. variation in solar availability and moisture levels (north- vs. south-facing This is the ®rst major study of clonal spread in redwoods using slopes) and variation in disturbance regime (®re more common in the upland molecular genetic techniques. sites vs. ¯ooding in the riparian sites). Sites were widely distributed across It has been suggested that resprouters tend to be shorter than the forest, were at least 500 m from each other, and were pure stands of reseeders (Midgley, 1996); this is evidently not true when con- redwood estimated to have been last cut 50±70 yr ago. sidering coast redwoods. Our primary goals were to evaluate On each site, up to 45 adjacent trees over 10 cm in diameter at breast the importance of clonal spread in second-growth redwood height (dbh) were mapped, tagged, measured for dbh, and sampled for DNA. TABLE 1. Site characteristics for Sequoia sempervirens stands in Jackson Demonstration State Forest, north coast of California, USA. Area Stand Mean stem Surrounding sampled No. stems density diameter (cm) forest canopy Site (m2) sampled (no. stems/m2) (SE) character Site moisture Solar availability Riparian R1 1000 37 0.037 41.4 (3.8) Continuous Hydric±Mesic Moderate±Low R2 2600 45 0.017 41.6 (2.3) Continuous Hydric±Mesic Moderate±Low R3 1550 42 0.027 41.6 (4.3) Continuous Hydric±Mesic Moderate±Low North-facing N1 1600 42 0.026 28.7 (2.0) Continuous Mesic Low N2 4000 31 0.008 54.6 (6.6) Continuous Mesic Low N3 1500 43 0.029 40.6 (3.8) Continuous Mesic Low South-facing S1 1000 38 0.038 44.7 (6.4) Patchy Mesic Moderate±High S2 1700 37 0.022 48.0 (2.5) Patchy Mesic Moderate±High S3 1000 41 0.041 43.9 (3.3) Patchy Mesic Moderate±High 1142 AMERICAN JOURNAL OF BOTANY [Vol.
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